February 13, 2014

Human admixture common in human history (Hellenthal et al. 2014)

A string of recent papers argued for admixture in human populations at time scales from the Middle Pleistocene to recent centuries. A new paper in Science makes the point convincingly for extensive admixture in humans over the last few thousand years. The authors include the creators of Chromopainter/fineStructure software; the new "Globetrotter" method appears to be a natural extension of that method that seemed to work wonderfully well except for the limitation of producing only a tree of the studied populations.

The paper has a companion website in which you can look up the admixture history of individual populations.

While reading this study, it is important to remember its limitations. Two are immediately obvious: (i) admixture events can only be detected for the last few thousand years, as this method depends on pattern of linkage disequilibrium which decays exponentially with time due to recombination, and (ii) detection of admixture seems to depend on the presence of maximally differentiated populations from the edges of the human geographical range; for example, the Japanese appear unadmixed even though they are clearly of dual Jomon/Yayoi ancestry. On the other hand, the method does detect the admixture present in the San at a similar time scale.

The case of Northwestern Europe appears especially striking as none of the populations from the region show evidence of admixture. This may be because the mixtures taking place there (e.g., between "Celts" and "Anglo-Saxons" in Great Britain) involved populations that were not strongly differentiated. Alternatively, population admixture history may have preceded the last few thousand years and is thus beyond the temporal scope of this method.

An exception to the rule that populations at the edges of the human range appear to be unadmixed are the Armenians who appear to be the only * between the Atlantic and Pacific in Figure 2D (shown at the beginning of this post). The companion site lists their status as "uncertain".

Other results are more questionable; for example, the authors assert that Sardinians are an admixed population with one side being "Egyptian-like" and the other "French-like" whereas the ancient DNA evidence as it stands would rather indicate that Sardinians are the best approximation of Neolithic Europeans currently in existence and so are more likely to (mostly) possess a gene pool that traces back to ~8-9 thousand years in Europe. It will be quite the surprise if so many Europeans from 5kya or earlier look like modern Sardinians and ancient Sardinians don't!

The analysis of Eastern Europe is particularly interesting as it documents three way admixture (Northern/Southern/NE Asian) in most populations but two way admixture (Northern/Southern) in Greeks, estimated at ~37%. The authors claim that this is related to the Slavs, which seems reasonable given the 1,054AD age estimate. On the other hand, according to the companion website, the southern element in Greeks is inferred to be Cypriot-like and it's far from clear that the pre-Slavic population of Greece was Cypriot-like or indeed represented by any of the populations in the authors' dataset.

The three-way admixture in much of eastern Europe is not particularly surprising as history furnishes ample evidence for groups of steppe origin in the region during historical times. Some bequeathed their both language and name (e.g., Magyars), others only their name (e.g., Bulgarians) on the local Europeans, but records indicate a widespread presence of "eastern" groups in Europe from the time of the Huns to that of the Ottomans. A study of late Antique eastern Europeans from the Baltic to the Aegean may help better document how the twin phenomena of the eastern invasions and the spread of the Slavs shaped the present-day genetic diversity of the region.

I suspect that a few ancient samples will be far more informative for understanding the recent history of our species than the most sophisticated modeling of modern populations. Nonetheless, it's great to have a new method that maximizes what can be learned about the past from the messy palimpsest of the present.Science 14 February 2014:
Vol. 343 no. 6172 pp. 747-751
DOI: 10.1126/science.1243518

A Genetic Atlas of Human Admixture History

Garrett Hellenthal et al.

Modern genetic data combined with appropriate statistical methods have the potential to contribute substantially to our understanding of human history. We have developed an approach that exploits the genomic structure of admixed populations to date and characterize historical mixture events at fine scales. We used this to produce an atlas of worldwide human admixture history, constructed by using genetic data alone and encompassing over 100 events occurring over the past 4000 years. We identified events whose dates and participants suggest they describe genetic impacts of the Mongol empire, Arab slave trade, Bantu expansion, first millennium CE migrations in Eastern Europe, and European colonialism, as well as unrecorded events, revealing admixture to be an almost universal force shaping human populations.

88 comments:

"A new paper in Science makes the point convincingly for extensive admixture in humans over the last few thousand years".

Admixture is almost exssential for survival. Unless a population is expanding into a previoulsy unexploited environment lack of external genetic contribution will lead to inbreeding depression, and usually extinction. Inbreeding is a major problem in the conseravtion of threatened species.

"(i) admixture events can only be detected for the last few thousand years"

But it is absolutely certain that admixture has been occurring since some sort of 'humans' first left Africa, in fact long before that.

"and (ii) detection of admixture seems to depend on the presence of maximally differentiated populations from the edges of the human geographical range"

Generally speaking populations at the geographic margins of any species' range are the most different from the average. Genes that flow through a population are least likely to reach the margins. Or at least reach any margins other than the one they originate in.

I think the Sardinian results make sense, in the context of their method. Don't Sardinians have some substantial "Basal Eurasian" admixture? If so, it would explain why they are construed as a mixture of Egyptian and French, as they would have substantial admixture with an "African-like" element. So it is correct from that angle, but it is just factually incorrect. They are probably the least admixed Europeans, on a temporal scale

This study is interesting and informative but like Dienekes said has limitations. The fact they cant detect Anglo Saxon-Insular Celtic mixing within just the last 1,500 years is terrible. It is evidence for what I have been saying for a while though, north-west Europeans mainly descend from the same group of Indo Europeans(Y DNA R1b L11) who conquered Otzi and Gok4 like people in west Europe.

The hair and eye color difference though between Insular Celts and west Germans is hard to explain. Insular Celts have about 10-15% red hair, west germans 3-5%, west Germans have over 60% light hair and close to 50% blonde hair, but Insular Celts have under 40% light hair and under 20% blonde hair, Insular Celts have over 80% light eyes but west Germans have less than 70%. If they share so much common ancestry going back only 4,000-5,000 years why are their hair and eye color percentages so different? It could be random selection and a little bit of different ancestry.

@ Dienekes : ;he case of Northwestern Europe appears especially striking as none of the populations from the region show evidence of admixture. This may be because the mixtures taking place there (e.g., between "Celts" and "Anglo-Saxons" in Great Britain) involved populations that were not strongly differentiated. Alternatively, population admixture history may have preceded the last few thousand years and is thus beyond the temporal scope of this method."

IMHO, it is definitely the former. At least with regard to the Anglo-Saxon migration to England, it was very recent, in the big scheme of things (as we all know)

@ Dienekes. "The analysis of Eastern Europe is particularly interesting as it documents three way admixture (Northern/Southern/NE Asian) in most populations " Yes but this is 2-4 %. barely significant. In fact, I would have thought it to be higher, Although it depends what they mean by "N-Et Asian". And this admixture would not be due to magyars, Bulgars or even Huns. Contrary to what many of the uniformed beleive, all these people came not from the far east (eg Mongoliga), but the Don-Volga - Kazakhstan region at farthest. So we should not expect to see "Asian" genetic signatures in these peoples, as they are ultimately 'west Eurasian'.

Also it is interesting that Romanians show far more 'soutner' component c.f. Hungarians, although both are placed at equal latitude.

I would be very sceptical for many of the results, especially for Greek-Slavic admixture. This study basically contradicts all previous studies both based on Y/mtDNA data as well as aDNA, which indicate a highly variable slavic ancestry in Greeks, not above 23%. It is interesting that Bulgarians (who look more slavic) in this study, have less ''Polish-like'' slavic ancestry than the Greeks. It is also unfortunate that other south slavic populations such as the Serbs and the Bosnians - characterised by an excess of Y-DNA haplogroup I were not included.What is also weird that Greeks have a significatn (6%) Scottish and English -like admixture.I would interpret this in two ways. 1) The Greek sample consists mainly of Northern Greeks (e.g. Macedonia, Thrace) which have both higher slavic ancestry compared to the rest of Greece and have historically been settled by Celts.2) As North Italians (always according to this study) have more than 30% ''scottish-british'ancestry, I would interpret this admixture in Greeks as a possible signal of Roman admixture.

Anyway, I think it is bad practice to try to make inferences about any large-scale migration by using such a limited sample (only 20 Greeks) and only two out of the many countries that were affected by it. In my opinion this study does not contradict in any way the assumption of all previous studies of a low slavic ancestry in Greeks.

You are right that Magyars came from the European/Central Asian border region and they were very probably dominantly West Eurasian/European to begin with. But Bulgarians and Huns originally came from much more far east. They simply got heavily mixed on the Steppe before they reached this region and even more later.

This characteristic difference between Romanians and Hungarian was already seen in Dienekes' Dodekad results that put Romanians very close to Bulgarians. But this is not strange at all. 1. Hungarians originally came from a region that had more North European like influence than the Balkan and later their main sources of heavy admixture were Germans and various Slavs (I'd even say the genes of the modern Hungarians are drawn more from Slavs and Germans that from the original Hungarians). 2. Romanians, who are the descendants of the Romanised Balkan population spiced up with some III. century northern refuges, moved North from the Balkan in the Medieval times (11th-14th century) and understandably have very close genetic ties to Bulgarians (they lived in the exact same area and have common genetic roots).

Terrible name and terrible conclusions. The study may have scientific flaws, but ADMIXTURE results, historical accounts, and linguistic accounts show very little difference between Celts and Germanics. End of story - moving on. You can't base hair colour and eye colour percentages in any sort of rational argument. These are autosomal traits which have been selected in and out of populations for various reasons. The higher rate of red hair in UK/Ireland may have to do with it being an island population. There were people already in Europe at the time of red hair mutations entering Europe to local populations who had dark hair and skin, therefore we cannot assume these mutations to be retained, although they have been to a degree.

I tried to understand the logic of the paper but I couldn't. What does it mean that Sardinians are Egyptian-like? How come Greeks are related to Poles, but not to Russians or other mediterranean peoples? How can Germans be related to the Finnish, but not to Poles? Reading the chart made my head hurt.

How many times do you need to be told that there was no Indo-European population replacement in Northern Europe. There was far more violence in Britain during the Neolithic than the Bronze Age.

British and Scandinavians are already close genetically to the British due to shared ancestry in the Mesolithic, Neolithic and up to the early Bronze Age. 20,000 Saxons entering a region of Britain with over 500k is not going to end in a slaughter or replacement. There are not 500k bodies.

It took 200 years for the North Germanics to become a 1/4 of the population of modern England. They were the dominant political and military force, not the dominant in terms of numbers. These British groups remained outside of Saxon and Angle influence until late into the 6th and sometimes the 7th centuries. They were later annexed and began mixing more later.

Lindsey and the whole East Midlands was significant in the number of British living between Saxon and Angle settlements. Bernicia was mixed from almost the get go.

The fact is that there was never a replacement in Britain. Not even during the Neolithic. Which is unusual for Europe. There are fewer remains showing violence from the Bronze Age, than the Neolithic. End of story.... Spare everyone the pseudo...

I see some weird African admix results in Northern Europe and also limitations in naming regions and identifying admix sources. Also, it is misleading to use present-day national names in Northern Europe, although it makes simpler to localize things. But it definitely leads to misunderstanding of the history. Of course this is a very common way to talk about ancestry, but also the most common way to avoid finding what really happened in our past.

"The fact they cant detect Anglo Saxon-Insular Celtic mixing within just the last 1,500 years is terrible."

If you agree that this actually occurred to any significant degree. Otherwise it is evidence for the opposite, that the English are not as Anglo-Saxon as they were led to believe. That most of them are not Anglo Saxon at all.

"It took 200 years for the North Germanics to become a 1/4 of the population of modern England. "

Where did you get this from? I don't believe it to be true. It is certainly not reflected in modern genetic differences between the Irish and the Brits. Autosomally these populations are almost completely overlapping with only very minor skewing in South eastern counties, and in the far far north.

@George D: You can't read these pop % too narrowly (as if excluding all other pops). If it shows Greeks as "part Polish and part Cypriot," we can read that as "part (something E European) and part (something Mediterranean/Mideast)."

Paper says as much. So the "25% (if memory serves) Orcadian/German/Austrian" in Kalash from 800-600 BC could really be a Greek or some other pop that's somehow related to West Europe (maybe just in a peripheral way). Behaves like the "Gedrosia %" in Orkney etcetera that nobody has even explained or delved into for some reason.

"It took 200 years for the North Germanics to become a 1/4 of the population of modern England."

West-Germanics.

Since English, Dutch and German are West-Germanic languages (Northgermanic = Icelandic, Norse, Swedish, Danish. Eastgermanic = Gothic), it must be "West-Germanic" people. At least those, who made the base for the English language. Even if Angles came from what is now Denmark, English is not a North-Germanic language.

George D. wrote"How can Germans be related to the Finnish, but not to Poles? Reading the chart made my head hurt."

Don't blame you. Especially if the Poles and the Lithuanians show admixtures with one another of around 50%. Equally curious is that there is supposedly no Polish admixture in Germany and no German admixture in Poland.

Since the program is seeing no admixture event in "Germany Austria" at all since 1300 BCE, it probably makes sense to ask when those admixtures shown for the Germans might have happened. I suppose they are earlier than 1300 BCE. And the largest admixture in Germany are from Finland, Spain and Lithuania -- but not vice versa. No German admixture is shown in any of those three.

I think one of the problems here is the measure of time. The other is sampling. In terms of 1000 years ago or more, the current political names are arbitrary. In East Germany, the backward projections with a larger sample would be I think very different.

Chad Rohlfsen said..."British and Scandinavians are already close genetically to the British due to shared ancestry in the Mesolithic, Neolithic and up to the early Bronze Age."

Yes, but there might have been admixing going on all along, but on a smaller scale than the program can make sense of. There were probably something Saxons in England a long time before the Saxons came.

It is really odd to see Norway represented twice in the data for England -- once as the source of an admixture of "(WestSicilian-like)" (5.1%) and only 2.9 for "(Welsh-like)" One does not think of the Norwegians as West Sicilian like - especially not when the program seems to date this admixture.

"I would be very sceptical for many of the results, especially for Greek-Slavic admixture. This study basically contradicts all previous studies both based on Y/mtDNA data as well as aDNA, which indicate a highly variable slavic ancestry in Greeks, not above 23%. It is interesting that Bulgarians (who look more slavic) in this study, have less ''Polish-like'' slavic ancestry than the Greeks. It is also unfortunate that other south slavic populations such as the Serbs and the Bosnians - characterised by an excess of Y-DNA haplogroup I were not included.What is also weird that Greeks have a significatn (6%) Scottish and English -like admixture."

It's quite possible that the elevated level of 'Polish-like' is due to persistent flow of of peoples from the Balkans. The classical Greeks recognized that they were an admixed people, and that much of their populations were derived from immigrants from the Balkans, many of whom originally spoke non-Greek languages such as Dacian/Thracian and later Albanian. The "Scottish and English-like admixture" in Greece may be related to historically-attested invasion by Celts, some of whom also settled in Anatolia.

What is really "off-the-wall" are their admixture results for the Pima, who show components of both Greek and Sardinian ancestry. In fact, the results presented in this paper show the Pima as having 5.6% Greek as opposed to only 4.6% Karitiana! Too bad this study didn't present results for populations in more northerly parts of North America.

@barak"The hair and eye color difference though between Insular Celts and west Germans is hard to explain."

It's easy to explain. If you add extra depigmentation to red-haired and green-eyed people you get blond hair and blue eyes.

So a simple explanation would be the Germans were originally similar to the insular Celts and then got extra hair and eye depigmentation that the insular Celts didn't, a larger influx of Indo-Europeans for example.

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"If they share so much common ancestry going back only 4,000-5,000 years why are their hair and eye color percentages so different?"

A simple explanation would be WHG and ANE are very closely related but the Indo-European part of ANE picked up some extra depigmentation genes from somewhere along the way so the blond to red cline is a proxy for IE percentage.

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@Chad

"It took 200 years for the North Germanics to become a 1/4 of the population of modern England."

The Danes are 1/2 R1b and 1/2 I1. The idea that the Anglo-Saxon marker in England is just I1 is wrong. Unless something has changed in the interim the Anglo-Saxons would have been 1/2 R1b and 1/2 I1 also.

So if I1 is c. 25% in England then that would make the Anglo-Saxon percentage c. 50%.

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R1b is more western than specifically Celtic and I1 is northern so the northwest is a bit of both.

"the southern element in Greeks is inferred to be Cypriot-like and it's far from clear that the pre-Slavic population of Greece was Cypriot-like or indeed represented by any of the populations in the authors' dataset."

As I have mentioned many times, we have to keep in mind that from archaeology, most of Italy and the Balkans all the way to the southern Pontic region were rather different to the rest of Europe after the initial Gravettian impact. After LGM, W and N and C Europe had the Magdalenian culture, Italy and the Balkans a different, Epi-Gravettian one - because of severe geographic barriers. But, there was absolutely no geographic barrier to Anatolia or the northernmost Levant. Thus, the simplest explanation is that just before the Neolithic, all those SE populations were very similar to each other, enhanced by intense sea trade, but different from the remainder of Europe.

The Greek northerly admixture probably goes partially back to IE times (from the western Pontic), and the "Slavic" admixture in the Balkans, in particular in the West, in reality is likely a conglomerate of Slavs and Eastern Germanic people, from historic accounts (hence also the unusually great body height, equaled only by Dutch people, and an occurrence of a very recent y-DNA I cluster).

"Also it is interesting that Romanians show far more 'soutner' component c.f. Hungarians, although both are placed at equal latitude."

Rob,

Hungarians usually place very close to Germans, Austrians, Poles, Czechs, and Slovenians in Admixture and PC studies. Romanians do not - see also my comment above.

As for the English, the problem is also that neither the Dutch, nor the northwestern Germans, nor the Danes are included. If the four included Germans/Austrians were from anywhere in central-western-southern Germany + Austria, they were no good match for the Anglo-Saxons, and the best admixture fit according to the analysis is with the Norwegians, and their contribution was little.

The result for the Tuscans is very strange! More specifically, it's the date that's puzzling: An Armenian-like element mixing with an English-like element around 942 AD, and definitely in Medieval times. The Armenian-like element may be tentatively brought in connection with Tyrrhenus' exodus, but what of that kind had happened in the middle ages?

Well, there have been Byzantine refugees in Tuscany, which were even one of the reasons for the beginning of the Renaissance there. But I'm surprised that the highest mixing coefficient is with Armenians and English rather than with Greeks and other Italians.

I agree that the case of Sardinia is interesting too. The dating of the admixture event completely rules out any connection with early neolithic events. From the date it can only be either Byzantine or Arabian, or both. The "Egyptian-like" influence is relatively weak, according to the analysis. The other, the "French-like" element has high admixture coefficients with the Spanish and the North Italians, would make some sense.

The classical Greeks recognized that they were an admixed people, and that much of their populations were derived from immigrants from the Balkans, many of whom originally spoke non-Greek languages such as Dacian/Thracian and later Albanian.

There is nothing in the ancient literature about Greeks being descended from immigrants from the Balkans, or indeed that "much of their populations" were descended from that source.

Interestingly, looking at the maps I see that side 1 and side 2 are both rather similar in Tuscans and North Italians, the main difference being the much later admixture date in the Tuscans.

As for the North Italians (from Bergamo I suppose), the 95% confidence interval of the admixture date allows for the possibility that it was the earlier population that was Cypriot-like, and that the later arriving Gauls brought in the Welsh-like element. However, the median value of the admixture date makes it more likely that the Cypriot-like element was brought in by „Roman“ colonists from southern Italy especially around the time when the north was granted the full Roman citizenship.

"The second process is explained through incentives, such as the Wergild outlined in the law code of Ine of Wessex which produced an incentive to become Anglo-Saxon or at least English speaking. The wergild of an Englishman was set at a value twice that of a Briton of similar wealth. However, some Britons could be very prosperous and own five hides of land, which gave thegn-like status, with a wergild of 600 shillings.Ine set down requirements to prove guilt or innocence, both for his English subjects and for his British subjects, who were termed 'foreigners/wealas' ('Welshmen'). The binary ethnic distinction that appears in Ine's Laws seems to be between ' Englisc/English ('us') and 'Wylisc/Welsh' ('them'). Since Ine's people self-identified as Saxons (West Saxons) this very early use of the word 'English' (unless it is a later introduction into the text) suggests that it was the use of a particular language, already recognised as a single language, and already called 'English', that was the crucial determinant in ethnic identity. This implies that in the early Anglo-Saxon period it was language use that was the key determination of ethnicity, and not whether you had "Germanic" ancestors."

"Oxygen and strontium isotope data in an early Anglo-Saxon cemetery at Wally Corner, Berinsfield in the Upper Thames Valley, Oxfordshire, found only 5.3% of the sample originating from Europe, supporting the hypothesis of acculturation. Furthermore they found that there was no change in this pattern over time, except amongst some females."

Also, there is confirmed migration to Britain from Scandinavia during the Bronze and Iron Age. Graves in Kent, during this period confirm it. Southern Britain also had a different diet from the late-Neolithic, on. There could've already been a genetic difference between southeastern Brits, and the rest of the island.

Trying to argue for Saxon aDNA, by the y-DNA is as useless as calling Irish 98% Siberian since they are 98% R y-DNA. Don't forget Belgic people migrated to Britain, pre-Rome. This would've introduced more s-28 and some s-21, if this wasn't already there in decent numbers. Migrations to Britain have happened for a long time. The Saxons were definitely not the first or last. To ascribe everything to them is just ridiculous.

eurologist said: the "Slavic" admixture in the Balkans, in particular in the West, in reality is likely a conglomerate of Slavs and Eastern Germanic people, from historic accounts (hence also the unusually great body height, equaled only by Dutch people, and an occurrence of a very recent y-DNA I cluster).

Frankly, I don't believe the elevated body height of West Balkan Slavs has anything to do with Germanic admixture. It would be strange if one trait, body height, was overtaken from Germanics and others like the cephalic index, nasal shape and pigmentation were not. Granted, in the presence of selection this may happen, but then selection may have fostered alleles for high body height that were already present in pre-Slavic/pre-Germanic times.

Anniemouse said: Otherwise it is evidence for the opposite, that the English are not as Anglo-Saxon as they were led to believe. That most of them are not Anglo Saxon at all.

Then why do they have plenty of R1b-U106 that correlates very well with Germanic admixture? And why are they inbetween the Germanic Dutch and the Celtic-descended Cornish according to the Dodecad K12b oracle? And why do they cluster with the Dutch and the Danes, according to the serological markers, and not with the Irish and Scottish? And why does all of this conform well to their West Germanic / non-Celtic language?

Overall, it seems that this paper is full of ambiguities...Interestingly, I even watched on the Greek news that based on this study the Polish are now considered as the cousin -nation of Greeks! Misinterpretation of scientific data by non-biologists is always funny :p A quick search in the literature on Greek autosomes greatly contradicts the results of this paper. Also, Dienekes, I know we basically have no idea of ancient Balkan autosomes, but wouldn't you expect slaves to have had an impact in the Greek gene pool and possibly introduce a more Northern component (if they had it)? I know most slaves were Greek, but there was a significant number of Thracian slaves (even introduced the worship of Mendis in Athens).A bit off topic, but an archaeologist friend in Athens tells me that there are so many bones found in excavations that sometimes they are even thrown away since there is nowhere to store them! How come nobody has attempted to extract DNA from these samples (I know that preservation is hard in hot countries)?

Haplogroups don't mean much. L-21 is still about as popular in England as S-21, while basically lacking in Germanic lands. L-21 is even with s-21 until you get to the southeast 1/3.

Look at the Irish, Niall of Nine Hostages haplogroup. One man who lived 1700 years ago, contributed to 21% of men in the region and 8% nationwide. So, technically it may only take 2-3 highly reproductive Saxon males to produce what we see today, without replacing genes, so let's not argue about haplogroups being people.

English plot between Irish and Dutch due to their location to the continent and receiving more geneflow from the continent during the Neolithic and metal ages. So, of course they are closer to the continent. People who live close, tend to be closely related in Europe. It's more about shared ancient ancestry than recent migrations. Saxons did not contribute the great deal of aDNA in Britain. Data doesn't match up. Not the DNA or the isotopes of graves, and grave goods.

I don't place much weight in Y chromosomes which tend to dominate in short periods of time, but if you assume (a false assumption in my opinion) that at the time of the Anglo Saxon invasion Britain was entirely "Irish-like insular Celt", then:

England is 14% I1Ireland is 7% I1Denmark 34% I1, 33% R1b

So at most 7% of English I1 would be available for calculations so 14% Anglosaxon men including R1b. Or 7%-14% of the combined population of men and women of Anglosaxon stock (depending on how many women migrated).

But then how do you explain the 28% I1 in Finland and 32% in Iceland. Not many Anglosaxons there, part of the natural European variation in I1. And then there is the matter that the gross chromosomes patterns are superficially very similar between the Netherlands (our nearest neighbour, 17% I1, 59% R1b) and England (14% I1, 67% R1b).

Not that you can rely on Y haplogroups anyway. But it is clear to me that the similarities in Y chromosomes between England, the Netherlands and Denmark are much older than the Anglo Saxon conquest. More Doggerland than the doggerel of Gildas and Bede, neither of who had first hand knowledge of the "genocide".

We actually have no idea what language most of England spoke at the time of the Roman conquest. There is some evidence (a phrase or two) that Cumberland spoke a Brythonic celtic. There are old English place names referencing the goddess Brigantia in England, Ireland, Scotland, Portugal, France, Austria and Hungary.

It has been argued that the Anglo-Saxon language took over in England because it was so similar to what they already spoke. But we have only to look at India to see how a language can come to dominate without significant genetic input.

Leonidas Davranoglu wrote: "Interestingly, I even watched on the Greek news that based on this study the Polish are now considered as the cousin -nation of Greeks! Misinterpretation of scientific data by non-biologists is always funny :pA quick search in the literature on Greek autosomes greatly contradicts the results of this paper.Also, Dienekes, I know we basically have no idea of ancient Balkan autosomes, but wouldn't you expect slaves to have had an impact in the Greek gene pool and possibly introduce a more Northern component (if they had it)? I know most slaves were Greek, but there was a significant number of Thracian slaves (even introduced the worship of Mendis in Athens).A bit off topic, but an archaeologist friend in Athens tells me that there are so many bones found in excavations that sometimes they are even thrown away since there is nowhere to store them! How come nobody has attempted to extract DNA from these samples (I know that preservation is hard in hot countries)?"

Mercenaries from Thrace as well as slaves from Thrace, Dacia, and (in the Byzantine era) Slavs were no doubt a major contributor to the populations of peninsular Greece. From "Dacia, Land of Transylvania, Cornerstone of Eastern Europe" by Ion Grumeza: "The Greek view of 'Dacian' was easily deduced from their definition of 'slave' as it was used in Athens. Indeed, in the 6th century BC , Dacian, Thracians, and other members of the Balkan tribes were sold in the slave markets within Greece under their name of origin." There were migrations from the Balkans to Greece - but not necessarily voluntary.

I'm saying people who assume that Celtic is R1b and I1 is Germanic are wrong. If the Danes are half and half and the Anglo-Saxons came from the same place then unless something happened in the interim the Anglo-Saxons would have been half and half also.

It is the assumption of I1 = Germanic that has led to all this silliness over the Anglo-Saxon invasion.

It's similar with people looking for very broad markers of the later Viking invasions and not finding them - lots of mostly the same DNA.

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"People who live close, tend to be closely related in Europe. It's more about shared ancient ancestry than recent migrations."

Yes, precisely.

"Saxons did not contribute the great deal of aDNA in Britain."

Or they contributed a great deal of aDNA in Britain but it was mostly the *same* aDNA.

You can see it on all the autosomal maps. The Atlantic coast / northwest were genetically mostly the same for a very long time but with tribal / cultural differences.

To me the currently known data clearly suggests that the English are a mixed people, of both Celtic and Anglo-Saxon ancestry. I don't understand why some people don't want to believe it. One reason may be that people want to have a clear, unambiguous identity, they want to be either Celtic or Saxon, but not a wishy-washy blend. Another reason may be the old aversion against Krauts. (Even though the Anglo-Saxons were partly from what is now Denmark.) As I wrote, neither the Dutch, nor a purely northwestern German sample, nor the Danes were included in this study. So the closest proxy of the Anglo-Saxons available in this study are probably the English themselves. Evidence from archeology and isotopes can never be as conclusive as the genetic properties and relations of the modern English. Occasional pre-Saxon influence of Scandinavians doesn't prove much. They didn't impose their language to the locals. And there was recently a story in the news about the presence of Egyptian immigrants in Roman Yorkshire. Does it mean much? I don't think so.

Frankly, I don't believe the elevated body height of West Balkan Slavs has anything to do with Germanic admixture. It would be strange if one trait, body height, was overtaken from Germanics and others like the cephalic index, nasal shape and pigmentation were not. Granted, in the presence of selection this may happen, but then selection may have fostered alleles for high body height that were already present in pre-Slavic/pre-Germanic times.

Simon,

You don't need selection, just a founder effect, which was definitely the case, as the star-like expansion and predominance of that particular y-DNA subgroup of I shows.

A lot of NW Balkans are also much more blonde, fair-skinned and blue-eyed than their Eastern neighbors at similar latitude. They definitely did not originate in the Southern Ukraine, based on the majority of their features. On the flip side, we know that people with such features lived in what is now W Poland, and by historical accounts spread SE and mixed with expanding Slavs, there, at the beginning of Slavic expansion.

As to the whole Polish admixture, don't forget that before Slavic expansion, what is now W and N Poland wasn't Slavic - neither culturally nor linguistically. And before the dominance of the people that brought the Baltic language to the Baltics, that region wasn't "Slavic," either. Finally, in both cases, the admixture from external sources did not leave a huge impact autosomally nor with respect to uniparental markers.

So, in conclusion, any "Polish" admixture has very little to do with "Slavic" admixture sensu strictu.

Can someone explain me how they date the admixture? In the case of the Greeks, obviously they don't compare modern vs ancient Greek specimens. So how do they know that 37% came during the middle ages? Do they compare Greek admixture to the to the Cypriot one?

If 37% of Northern admixture came during the Middle Ages, then the Ancient Greeks were purely Med and had almost no Northern admixture. One would expect that from the Minoans, but not the Greeks who presumably were more heterogeneous.

NY Times article."Dr. Myers and Dr. Hellenthal said that they hoped historians would find their work useful, but that they had not collaborated with historians."

“In some sense we don’t want to talk to historians,” Dr. Falush said. “There’s a great virtue in being objective: You put the data in and get the history out. We do think this is a way of reconstructing history by just using DNA.”

The above is a nice scientific summary although I dont agree with all of it.

In the end it is the autosomal situation that makes my mind up. The English are not different from the Irish despite Y chromosome patterns. There is no significant skewing of the English towards the Angles or Saxons. What overlap appears in genetic maps is what you would expect from a similar neighbour. Not a major genetic contributor.

I'm surprised no one has commented on the admix result for the Maya. Unlike the Pima, the Mayans show components from all over Eurasia, including the Caucasus and the Indian subcontinent. South American Native American populations such as Karitiana show no such links to Eurasian populations. Maybe this should not be surprising because various sources have indicated that Meso-American populations show a more East Asian component. Apparently, they also show more of a West Eurasian component not due to recent Iberian admixture. I was looking at a Mexican ancestry message board a while back and the number of Y DNA types thought of as uncharacteristic of Native American such as various subtypes of YHG N show up.

"There is nothing in the ancient literature about Greeks being descended from immigrants from the Balkans, or indeed that "much of their populations" were descended from that source."

It depends on how you interpret the Dorian Invasion (if indeed it existed), and assumptions about who the Dorians were and where they came from. One theory is that they were a non-Greek speaking people from farther north in the Balkans who became hellenized but were set apart from other Greeks by their distinctive dialect, which may have preserved elements of their original language. Interestingly, they wound up in the Peloponnese as did the later Slavic tribes such as the Ezeritai and the Melingoi who immigrated to Greece in the 7th century. Other Greeks certainly viewed the Dorians as different, and the root cause may have been their non-Greek origin.

Yes, I am aware of the controversy over Fallmerayer's theories, but I don't believe that Greece was ever depopulated and the original population replaced at any point during the Dark Ages, nor the claim the Slavs in Peloponnese were exterminated by the returning Byzantines. Historical evidence appears to indicate that Slavs who had settled in Greece were rapidly hellenized and assimilated following their submission to the Byzantines.

Some words found in Greek do hint at origins in other languages. For instance, although the Greek word for 'water' in the official Swadesh list is nero, the word particle for 'water' in the word for 'river' (potamos) is -am-, which resembles Dacian/Thracian apa or aba, except for the alternative consonant shift of -kw- --> -m- instead of -b-/-p-. This parallels the independent shift ahwa --> ama observed in the Cherokee word for 'water' as a result of its split from Northern Iroquoian languages. The word for 'sea' in Greek, thalassa, has baffled philologists. There is a possible analogue in Basque: itsaso for ’sea'. Note the ‘-aso’ word particle, which resembles '-assa’ in the Greek equivalent. The thal which replaces its- may be derived from tar through the phonetic shift ’t —> th’ and ‘r —> l’.

Haplogroup I1 is common in Germanics, but it doesn't correlate with Germanic influence as well as R1b-U106 does, presumably because it was already present and spreading before there were any Germanics in existence. For instance, afaik it's the second most common haplogroup in Scotland, after R1b.

(On a side note: Annimouse's observation that I1 in Iceland contradicts the assumption of an Anglo-Saxon origin of I1 in England is wrong; nobody suggested that I1 was exclusively Anglo-Saxon; instead some suggest that it is Germanic. And Iceland was definitely settled by Germanic Vikings, so no contradiction here.)

I wouldn't overestimate the importance of y-haplogroup frequencies either, but saying that they don't matter at all is clearly an exaggeration. And it's not reasonable to take strong founder effects and drift as the default assumption.

The modern Danes have about equal amounts of R1b and I1, but the Angles, Saxons and Jutes were not Danes, they spoke West- not North Germanic dialects. Linguistically English is closer to Dutch and Frisian than to either Danish and standard German. (Btw another reason why the anti-Kraut reflex is misguided here.)

If mere geographical proximity was the reason for the deviation of the English from their Insular Celtic neighbours, then they ought to deviate in the direction of the Belgians and northeastern French, not the Dutch and Danes.

Place names, tribal names and personal names of pre-Roman England show no difference to other Celtic regions. There is no evidence of a Germanic tendency there.

I agree on the point that genetically the Anglo-Saxons were basically northwestern Europeans too, and hence the difference to Celtic northwestern Europeans was not huge. I also think it's impossible to segregate Celtic and Anglo-Saxon influence neatly on the basis of phenotypes. There have been Early European Farmers and Dinaroid Bell Beaker folks in Jutland too, the latter's impact can still be seen here and there, particularly in northern Friesland for example. And think of the Tollund man; doesn't he look rather „Celtic“ with his narrow face, sloping forehead and convex nose? Yet he was a close relative of the Anglo-Saxons.

@ Eurologist, I agree that some Slovenians look strikingly German or Swiss, i.e. Central European (while others look distinctly Slavic), but this may be due to older expansions from Central Europe, like from the Hallstatt culture, rather than from the migration period. Actually according to Coon's old map on body height, the tallest Southern Slavs are from Bosnia and Montenegro, from rather secluded areas , that is. What Germanic influence did we have there? Just the Visigoths marching through.As for the Polish admixture: The study we are discussing here seems to suggest that the Poles are about 2/3 Slavic, 1/3 pre-Slavic.

@ Constantine, the dates are inferred from the average length of the genetic segments that can be ascribed to either of the source populations, the shorter the segments, the older the admixture.

The southeastern 1/3 of England was most likely Belgic speaking. I'd put Saxon admixture at maybe 10-30%, increasing to the south and east. 30% might be generous too.

Chad,

At what time? Archaeology puts Germanic-type houses in the E England context before post-Roman invasions. It looks like the Roman award system and NW Germanic (W Frisian/ N Frisian, Saxon, in that order) followers conquered early and quickly.

I had heard that Belgian speakers were brought in rather late for their writing capabilities - which introduced some Germanic oddities into written English (silent k, silent gh instead of voiced ch, etc.).

"But it is clear to me that the similarities in Y chromosomes between England, the Netherlands and Denmark are much older than the Anglo Saxon conquest."

I agree that the DNA distribution is older and the problem here stems from assumptions that specific y dna map cleanly onto specific groups.

.

"More Doggerland than the doggerel of Gildas and Bede, neither of who had first hand knowledge of the "genocide"."

However none of these mis-assumptions are anything to do with Bede's history which as far as I can tell is being pretty much totally vindicated by genetics as long as there isn't an assumption that specific y dna maps cleanly onto specific historical groups.

And there was never any need to assume a violent genocide from Bede's history.

Bede's version goes:1. Roman's leave.2. Saxon pirate attacks along the east coast.3. Romano-British cede parts of the east coast to Saxons in exchange for fighting off the pirates (just like the Normans in Normandy): creation of the "Saxon Shore."4. Mass migration from the source regions settling along the east coast.5. Big war.6. Stalemate with an east-west division of the country between Saxon and Romano-British.7. Piecemeal conquest of the west of the country over the following centuries.

Looking back at stage (2) if a population is being raided by people they can't defend against because the attackers are more mobile than them e.g. sea-borne or horse-borne, then the target population will **move away**.

The Romano-British along the east coast would have depopulated because of the pirate raids before the main Saxon incursion in the same way the med coast depopulated during the time of the Arab slave raids.

Once the Saxons settled they lost their raider advantage and the conflict became more balanced leading to a stalemate (exactly the same as happened later with the Danes).

There's no need for violent genocide or apartheid states. All you need is a lot of raiding before the main invasion / migration leading to depopulation.

#

"Frankly, I don't believe the elevated body height of West Balkan Slavs has anything to do with Germanic admixture."

False correlation maybe.

Maybe the height comes from the pre-IE HG population that only survived as a large percentage in some parts of the Balkans and the parts of northern Europe that *became* Germanic later.

To me another good argument in favour of a considerable Anglo-Saxon impact in at least parts of England is the striking phonological similarity between the Low German dialects in Schleswig-Holstein and surroundings and the English accents in large parts of England, except for the west and north. Linguistically, Frisian may be the closest relative of English. But as far as the mere sound of the language is concerned, I dare saying that the Low German of northernmost Germany is the closest. Of course I'm not talking now of American English or other derived variants, but of RP aka BBC English and of similar accents.

Take this old recording of the old Low German dialect of Hamburg as an example:

http://www.staff.uni-marburg.de/~naeser/ld011k.mp3

The melody of the language, the way the vowels are diphthongicised and the way the t is pronounced are just some of the similar features.

I think it's not easy for a non-native speaker to imitate this pronunciation. And often, when a language is imposed on a substrate language by elite dominance, the substrate exerts some influence on the phonology of the dominant language. I think in England we see exactly this: The southwestern accents sound very different, somewhat similar to Irish English, and I think that's because of the stronger Celtic substrate there.

Gary, actually Greece *was* depopulated severely from c. 700AD. Several massive land surveys by Anglo-American teams confirm thisOnly a handful of towns survived , but it was the result of general roman trouble and not due to SlavsIndeed, much of Greece *was * repopulate a by Slavs who gradually became Greek speaking, augmented by Armenians, and Greek colonists from Asia Minor etcWhether this sits well with Greeks is beyond the pointBut surely admixture is a good thing :)

It depends on how you interpret the Dorian Invasion (if indeed it existed), and assumptions about who the Dorians were and where they came from. One theory is that they were a non-Greek speaking people from farther north in the Balkans who became hellenized but were set apart from other Greeks by their distinctive dialect, which may have preserved elements of their original language.

You claimed that the _classical Greeks_ recognized that many of their ancestors came from the Balkans. That is quite different than saying that _modern theories_ might place the Dorians' origin to that source.

The classical Greeks never derived the Dorians from the Balkans, or indeed any substantial population element from that area. So, whatever argument one can make for Balkan->Greece migration in antiquity, it's best to leave the classical Greeks and what they thought out of it.

All tribes in the SE 1/3 pre-Rome are accepted as Belgic Celts who migrated to Britain starting in 250BCE. The material culture is the same as the continent, exactly! Romans were after Belgic people who joined kin in Britain, before the conquest.

Here's a couple of plots. The British are as close to the Irish and Belgians as they are the Dutch. They do not sit on top of the Dutch, which you might expect with your wipe-out theories. I am sorry but there is no way that 95% of Brits were eliminated anywhere. Scots are closer to the Dutch than the most southern Brits.

As for Bede, no one takes his account seriously. I already explained and gave references for the continuation of British culture next to Saxons. There was never a whole give up of the East coast. There are British settlements on the east coast up to the 7th Century. There is no 'Anglo-Saxon' identity until we get to the end of the Danelaw period. There are at least 35 groups in the old tribal hideage. Look it up. There is no single identity anywhere.

Lastly on the language, English is not like those Germanic languages. Germanic philologists agree, almost completely that the syntactic features of English is not like other German languages because of a large acculturation event of Celtic and Latin speakers who were native to Britain, switching to Old English. Brythonic had a huge effect on English. The 'Celtic hypothesis' is widely accepted. I've already provided links that show Britons outnumbered 'Saxons' by 4-1 in 700AD. Bronze and Iron age Celtic would've become dominant the same way, an elite take over with no replacement. Nothing changes with the Saxons. There was no unifying Anglo-Saxon language until the 9th Century.

Ward-Perkins, ‘Why did the Anglo-Saxons’, 258, suggested that the successful native resistance of local, militarised tribal societies to the invaders may perhaps account for the fact of the slow progress of Anglo-Saxonisation as opposed to the sweeping conquest of Gaul by the Franks.

Toon, T.E. (1983) The Politics of Early Old English Sound Change. New York

Lastly is the issue of elite names. Look at all of the 'Saxon dynasties', that have Celtic personal names. Cerdic, Caedwallen, and Ceawlin, just to name 3. There is more of a Briton etymology to names in Penda's line in Mercia as well.

Personally, I think that it is much more likely that English on average are 50% Briton, 25% Belgic/Gaul, and 25% Germanic. Germanic will be higher in East Anglia and York though. No where near a replacement number, of course. I would also safely bet that the south coast is more native than lands to the north and east.

Gary Moore wrote:"It depends on how you interpret the Dorian Invasion"

I do remember those theories ("this particular Dorian sub-tribe or that particular sub-tribe were such-and-such because of this word or this tribal name or this place name") but they do seem kinda hard to swallow. West Greek (Doric + NW Greek) covered by far the biggest area in mainland Greece (from the southern Peloponnese all the way to the Acroceraunian, excluding Arcadia; and perhaps even Upper and/or Lower Macedonia but that depends on your perspective :)). What exactly is the proposed mechanism there?

"but I don't believe that Greece was ever depopulated and the original population replaced at any point during the Dark Ages, nor the claim the Slavs in Peloponnese were exterminated by the returning Byzantines. Historical evidence appears to indicate that Slavs who had settled in Greece were rapidly hellenized and assimilated following their submission to the Byzantines."

That does seem to be the case (though decline in population and migrations obviously occurred as you say; interestingly enough, around the same period there is evidence of depopulation in the area of Poland and there are various schools of thought about what exactly and to what extent happened there with the similar back-and-forth that plagues these issues everywhere) and it's a nice confirmation of what the majority of historians and archaeologists who didn't follow unfounded extremes like "total depopulation" or "swamping" had arrived at.

"For instance, although the Greek word for 'water' in the official Swadesh list is nero"

Do keep in mind that "nero" is metonymical, though, and that "hydor" is the word you should use. I'm also not quite sure why you analyze "potamos" that way (or why it should be a derivative of "water"). I looked at both Chantraine and Beekes and nothing like that appears in either; they prefer words that have to do with movement but Beekes thinks it might be non-IE 'pre-Greek.' The -ss- and -tt- words are well-known to be part of a pre-Greek sub-stratum so under some pre-Indo-European Vasconic substratum hypothesis, that could work (if the changes are correct; I have no clue, I'm taking your word for it).

eurologist wrote:"A lot of NW Balkans are also much more blonde, fair-skinned and blue-eyed than their Eastern neighbors at similar latitude"

How 'much more,' though, can you quantify it?

"So, in conclusion, any "Polish" admixture has very little to do with "Slavic" admixture sensu strictu."

Sensu stricto no, but how can you tell which original group ("Goths" or "Slavs") was responsible for that specific thing and how do you discount the possibility of pre-Völkerwanderung sources based on the little we know so far? FWIW, elevated height seems to be a feature of the whole Balkans when compared with surrounding populations (south Albania and Bulgaria excepted perhaps which seems hard to explain purely under admixture scenarios; or perhaps it's a matter of nutrition). The others just don't tend to be as tall as the Dinaric populations.

Actually, parts of your theory are compelling to me but it'd be much better if the study had included Ukrainian samples, at any rate.

"Indeed, much of Greece *was * repopulated a by Slavs who gradually became Greek speaking, augmented by Armenians, and Greek colonists from Asia Minor etc".

Then how come R1a and I2 levels are so low in Greece? Most of the re-population happened by Greeks from Magna Grecia, who brought many loanwords from Latin. This was organised by the Byzantine rulers of that time. In fact the common patronymic "-opoulos" ending of many Greek surnames (mostly associated with the Peloponese region) is of Latin origin (pullus) and it is thought to originate during that period.Regardless, I agree, admixture is a good thing.

There is no way how the peculiar sound of (some) of the more eastern to central dialects of England could be a legacy of ancient Belgic. First of all the area settled by the Belgae was also the area of the strongest Roman/Latin influence. And then it's noteworthy that just a few miles west of Hamburg, on the mouth of the Weser, the dialect sounds nothing like English:http://www.staff.uni-marburg.de/~naeser/ld010.mp3

Neither does Westphalian:http://www.staff.uni-marburg.de/~naeser/237nr.mp3

Nor Low Frankish:http://www.staff.uni-marburg.de/~naeser/ld231k.mp3That's already essentially Dutch. Dutch and Flemish don't have that peculiar English sound either. It's confined to Schleswig-Holstein and nearby Hamburg, the southern part of the traditional homeland of the Anglo-Saxons.

What is interesting to me is for "East Sicily", going by the dates for estimated admixture events, the original population was proposed to have been 654 BCE and consisted of both British and Irish-like people as well as Northeast European, while the later migration was estimated around 1474 CE and consisted of both Northern Italian-like people and a host of East Mediterranean (Levantine, Arabian, etc) and North African groups as well as Greek. This is inconsistent with the historical record of Phoenician and Greek colonization, as I'd expect the northern input to have been mostly Norman and would have come way later than the influence from the Near East.

It's hard for me to believe the original population of Sicily was Irish-like. I always suspected the original population would have been Near Eastern in character and acquired northern elements more recently with all of the Lombard, Norman, French and Spanish influences that came during the Middle Ages.

Dr. Rob said:'Gary, actually Greece *was* depopulated severely from c. 700AD. Several massive land surveys by Anglo-American teams confirm thisOnly a handful of towns survived , but it was the result of general roman trouble and not due to SlavsIndeed, much of Greece *was * repopulate a by Slavs'

A research which I have come across claims quite the opposite. That recent excavations have shown that Greece proper was not depopulated from 700 A.D. and that there are signs that there were reasonably big cities with high economic activity.

The truth is that when it comes to the state of Greece proper or its demographics during this time we are second guessing with a scarcity of information. We don't know much about the Slavs in Greece either. The very few medieval sources who mention anything about Slavs in Greece (e.g. the chronicle of Monemvasia) are not very reliable and include texts which are clearly fantasy. Many historians dispute the reliability of the source and claim it had a certain agenda. I would rather rely more on genetics and anthropology on this one.

Dienekes said:>>>"The classical Greeks never derived the Dorians from the Balkans, or indeed any substantial population element from that area. So, whatever argument one can make for Balkan->Greece migration in antiquity, it's best to leave the classical Greeks and what they thought out of it."

People in some isolated areas in Crete where we know Dorians settled show North European elements. Perhaps the Dorians brought more northern admixture in Greece. Dorians never settled in Cyprus. It would be interesting to compare Sfakians and Cypriots and see if Sfakians have much more distinct East European origin compared to Cypriots. Perhaps it would say something about the origin of the Dorians.

This parallels the independent shift ahwa --> ama observed in the Cherokee word for 'water' as a result of its split from Northern Iroquoian languages.

Fascinating, are you aware that the Irish Gaelic word for river is "Abhainn", pronounced "OW-an", and that the Welsh word for river gave us "Avon" in Stratford-on-avon...Makes is seem like the Iroquoian word is similar to the Celtic word?

ABBD said:'Then how come R1a and I2 levels are so low in Greece? Most of the re-population happened by Greeks from Magna Grecia, who brought many loanwords from Latin. This was organised by the Byzantine rulers of that time'

Greece has higher levels of E-V13 than any region of Magna Graecia. Especially the mainland, but Greece in general as well. A population with lower levels of E-V13 can not populate a place and increase the level that haplogroup. And surely the Slavs did not have anything to to with it. I also doubt all R1a in Greece came with the Slavs, but that's another matter.

Genetically these migrations could only have occurred on a smaller scale than some claim.

Please compare Celtic words with other Indo-European words before comparing them with words from the languages of indigenous tribes of America, Australia, Africa, or whatever distant place you may like.

In the case of words like abhainn or avon, I would suggest considering Latin aqua ("water") or Iranic âb, etc. ("water; river").

The following data are from a table on the Wikipedia page for "Iranian languages":

"Here's a couple of plots. The British are as close to the Irish and Belgians as they are the Dutch. They do not sit on top of the Dutch, which you might expect with your wipe-out theories."

I don't have wipe-out theories. That's your straw man. I have [people move away from places where they can be raided with impunity] theories.

Sitting halfway between Irish and Dutch is exactly what you'd expect from Bede. A first stage involving a wholesale Anglo-Saxon takeover of the east coast followed by a second stage of slow piecemeal elite conquest resulting in an east to west cline with a roughly 50/50 balance overall: 90% A-S on the east coast, 90% pre A-S on the west coast, 50/50 in the middle.

Again, exactly as you'd expect from Bede.

.

"Scots are closer to the Dutch than the most southern Brits."

Scots along the east coast were Anglo-Saxons too.

.

"As for Bede, no one takes his account seriously."

Well they should as the genetics support him.

.

"There is no single identity anywhere."

Who said there was? Bede, again, talks very clearly about there being lots of separate groups involved in the invasion including Angles, Saxons, Jutes (Geats) and Frisians and sub-groups within them e.g. West Saxons and East Saxons etc.

"Lastly is the issue of elite names. Look at all of the 'Saxon dynasties', that have Celtic personal names. Cerdic, Caedwallen, and Ceawlin, just to name 3."

Yes the royal lines start off with Saxon names in the east and then as the Saxon kingdoms expand west they often take on Briton names almost like the population was getting more 50/50 as it expanded west - which it was.

Personally I don't think I1 is Germanic exactly. I think I1 is northern WHG that mixed with the northern branch of R1b expansion but it acts as a kind of Barium meal for Germanic ancestry and you can quite plainly see the east-west cline in Britain: majority A-S in the east, minority in the west, 50/50 in the middle.

Alogo wrote - "Do keep in mind that "nero" is metonymical, though, and that "hydor" is the word you should use. I'm also not quite sure why you analyze "potamos" that way (or why it should be a derivative of "water"). I looked at both Chantraine and Beekes and nothing like that appears in either; they prefer words that have to do with movement but Beekes thinks it might be non-IE 'pre-Greek.' The -ss- and -tt- words are well-known to be part of a pre-Greek sub-stratum so under some pre-Indo-European Vasconic substratum hypothesis, that could work (if the changes are correct; I have no clue, I'm taking your word for it)."

The pot- component does refer to "flowing" in Proto Indo-European. Presumably, the word means "running water". The -am(a)- fits the general pattern of phonetic shifts of -kw- to labial consonants (b/(m/(p/(v in Indo-European languages:

Latin aquaDacian/Thracian aba/apaPersian ab(a)Kurdish av(a)

A language with the shift -kw- --> (-m- can be inferred, and it appears that such a language did exist and was absorbed into ancient Greek without being attested, but its legacy survives a word particle.

The language of the Pelasgians is unknown. Although it has been commonly assumed to have been "pre Indo-European", it may have belonged to another branch of IE.

Also: "I do remember those theories ("this particular Dorian sub-tribe or that particular sub-tribe were such-and-such because of this word or this tribal name or this place name") but they do seem kinda hard to swallow. West Greek (Doric + NW Greek) covered by far the biggest area in mainland Greece (from the southern Peloponnese all the way to the Acroceraunian, excluding Arcadia; and perhaps even Upper and/or Lower Macedonia but that depends on your perspective :)). What exactly is the proposed mechanism there?"

There are competing theories about the collapse of Mycenaean civilization. In one model, Doric was simply the language of the peasants, which was more archaic than the language of the palace culture. On the other hand, the Dorians may have been an admixed population from the border areas of northwest Greece who, being on the fringe of Greek culture, may have spoken a more archaic version of their mother tongue. The legend of the "Descent of the Heracleidae" associated with the "Dorian invasion" could have a grain of historicity: the theme of an exiled ruling clan going into exile and returning with a mercenary army to reclaim their throne is a familiar one. Of course, when this happens, things have a habit of getting out of hand...

Ebizur wrote: "Please compare Celtic words with other Indo-European words before comparing them with words from the languages of indigenous tribes of America, Australia, Africa, or whatever distant place you may like."

Iroquoian languages are probably related to Indo-European, and the farther east the IE language, the more they look alike. (Some Iroquoian words bear a remarkable resemblance to their equivalents in Indo-Iranian languages.) It looks like Indo-European probably began with a migration out of Siberia by a people who spoke a language related to the Iroquoian languages of North America, and who later admixed with other peoples while moving through Central Asia and the Pontic Steppes before entering Europe. Some prominent linguists have long maintained that many Eurasian languages are related to the Dene languages of North America, and it now appears that IE may be related to a North American language family as well.

I think it probably needs refining so I don't take it that seriously yet but taking it at face value just for fun and assuming:

1) These mixtures don't relate to the actual populations but shared ancestral components,

2) The earliest expansions were waves of very low population density foragers coming out of particular regions in a particular time sequence with the earlier waves being mostly but not completely replaced by later waves,

then very widely dispersed components that don't seem to make sense would most likely be remnants of a submerged wave - like a high tide marker.

If so the oddest ones should always be in very inhospitable terrain.

So say the first three steps of the sequence were:1) Out of the Tropics2) Out of Africa3) Out of India

The Lithuanian-Hadza link might represent remnants of the stage (2) expansion that survived in some deep swamps around the Baltic and in the Hadza desert.

The Scottish-Kalash example might represent remnants of the stage (3) expansion that survived up remote mountains.

While elsewhere both those components were mostly replaced by later expansions.

Here's a couple of plots. The British are as close to the Irish and Belgians as they are the Dutch. They do not sit on top of the Dutch, which you might expect with your wipe-out theories. I am sorry but there is no way that 95% of Brits were eliminated anywhere. Scots are closer to the Dutch than the most southern Brits.

Don't count me to the wipe-out theorists, I've never claimed anything other than that the English are mixed. The Dodecad results made me think of 50-50, but the exact figure is open to debate, and that will only be settled with ancient DNA.

As for your plots, the first plot shows a significant overlap of the UK with Scandinavia. I'm pretty sure that this is partly due to the Anglo-Saxon influence, but since the sample is not exclusively English, other connections to northern Europe are probably also at play. Your other plot rather vindicates my position; GB deviates from Ireland more into a Dutch/Danish/German direction than into a French/Belgian one, where the Belgae would have drawn it to. It's better discernible in this map:http://i57.photobucket.com/albums/g240/simon_w1/nature07331-f22_zps018ead64.jpg

The bulk of the British who deviate from their island are drawn right onto the North Sea, into an eastern direction, and not towards the northern French/Belgian area, although some indeed are.

There is no 'Anglo-Saxon' identity until we get to the end of the Danelaw period. There are at least 35 groups in the old tribal hideage. Look it up. There is no single identity anywhere.

Of course there were no Anglo-Saxons on the continent. I used that term as a simplification when referring to Angles, Saxons and Jutes. (And there were also other Germanic tribes involved, like the Frisians, even Swabians as evidenced by the placename Swaffham.) I think it's not a big issue to collectively refer to them as Anglo-Saxons, since they were similar in origin, language and culture.

With reference to the language, I've read about some possible grammatical influence of the Celtic substrate too, so I wouldn't rule it out. But then there were also centuries of linguistic „drift“, modern English obviously is no longer the same as Old English, and there was a huge Middle French influence from the Normans, mostly in the vocabulary. Along the same line Low German received a lot of input from Frankish and High German, which made it deviate from Old Saxon, too. So while both English and Low German had the same Old Saxon root, they deviated from each other because of the separation and different external stimuli. Within West Germanic there once was a North Sea Germanic subgroup that included the Angles, Saxons, Jutes, Frisians and some other tribes. The Frisian language has preserved the North Sea Germanic features better than English and Low German, which might explain why it's considered the closest relative of English. The theory that English is derived from ancient Belgic is about as silly as the related theory that Alsatian is derived from Gaulish (which once had a proponent too).

Check out these guys from western Holstein:http://www.staff.uni-marburg.de/~naeser/ld185k.mp3

The first one sounds like an Englishman who's speaking German with an accent.

Lastly is the issue of elite names. Look at all of the 'Saxon dynasties', that have Celtic personal names. Cerdic, Caedwallen, and Ceawlin, just to name 3. There is more of a Briton etymology to names in Penda's line in Mercia as well.

Others had Germanic names, like Raedwald, Aethelberht, Egbert, Edwin, to name but a few.

@pconroy, Orcadian % in Kalash 800-600 BC is IMO not Macedonian or anything to do with Alexander (too early by several centuries).

Could be from Medean influence in Central Asia (cities like Mundigak). Assyrians deported people to some Medean cities as imperial policy.

Medes were founding culture that another population (Persians) apparently adopted for their own empire. For Greeks, "Persian" culture was actually "Mede" culture. Rules like Arta (always speaking the truth), etc, Medes were apparently the "rough older brothers" of the Persians,

Also might be related to Levite R1a. Not necessarily exactly same pops as those that mixed with Kalash though (who knows).

The analysis says that around 1474 AD (1054 – 1698 AD), a West Sicilian-like population admixed with about 10% Egyptian-like people. Perhaps this may be from the Arabic invasion, though it looks a bit inconclusive (highest mixing coefficient with South Italians). The West Sicilian-like local population that was present before that admixture event (and which gave 90% to modern East Sicilians) had complex origins, with relations to western Europe, Greece, Italy, the Near East and the Caucasus.

The other admixture event, according to the study, was around 654 BC (with a very wide confidence interval, 6954 BC – 186 AD) and involved sub-Saharan and northern European populations.

Gary Moore said...I was looking at a Mexican ancestry message board a while back and the number of Y DNA types thought of as uncharacteristic of Native American such as various subtypes of YHG N show up.

Though there is the fact that Scandinavia and Russia have injected small pulses of religious dissenters into Mexico in comfortably post-Columbian times, this sounds unlikely to me. Could you please share the link?

Low-German is old-German and the original German. It was spoken by the tribes you mentioned. Frankish is franconian german and is basically west-central german along the rhine river.High-German is Bavarian, last tribe to become germans and bavarians colonised austrian. and Austria formed in 998AD from bavarian blood.Conclusion. The English had origianl old-German from Saxons, angles, jutes, frisians and others.The dutch also are a dialect of old-german

Low-German is old-German and the original German. It was spoken by the tribes you mentioned. Frankish is franconian german and is basically west-central german along the rhine river.High-German is Bavarian, last tribe to become germans and bavarians colonised austrian. and Austria formed in 998AD from bavarian blood.Conclusion. The English had origianl old-German from Saxons, angles, jutes, frisians and others.The dutch also are a dialect of old-german

I agree with your comment that comparing Cretans and Cypriots can give us some important clues regarding the origins of the Dorians and it is a pity Cretans are not included in this study. Generally I find it unbelievable that Cretans are ignored in practically all admixture studies of the Eastern Mediterranean region (see Behart et al., Haber et al, etc.).

Anyhow, a few months back I also proposed that Dorians may be an important source of the Northern European admixture among modern Greeks (http://www.eupedia.com/forum/threads/29183-What-can-autosomal-calculators-tell-us-about-ancient-Greek-admixture). After studying the autosomal amixture data of a few Cretans and comparing them with that of Greek Cypriots, they are strikingly similar, with Cretans having only slightly higher Northern European admixture. Therefore, I would suggest that the vast majority of the Northern admixture in modern Greeks came from the Slavs and to a lesser extent from Germano-Celtic tribes. This is also supported by the findings of the current study.

Alexandros said: Therefore, I would suggest that the vast majority of the Northern admixture in modern Greeks came from the Slavs and to a lesser extent from Germano-Celtic tribes

That at least some is "Migration Period" and afterwards is a given at this point (well, always was) but Cretans aren't necessarily a good stand-in for mainland Greeks and both Cretans and Cypriots have been subsequently affected by East/Southeast sources. Whether it was a "majority" or a "large minority" or...seems unanswerable at this point.

I wouldn't have formulated it exactly like this, but true, the High German language shift apparently started somewhere in the South; it was also performed most completely in the southernmost dialects; then there is a broad belt of middle German dialects which followed it partly, and then in the north, Low German, Frisian and Dutch didn't perform it at all, which makes them closer to old West Germanic.

I am not sure if the N.European features sometimes found among the people of Sfakia in Crete can be attributed to the Dorians. That part of Crete has been historically linked to an Eteocretan population, possibly pre-indoeuropean hunter gatherers, which could partly explain their different phenotype. As far as I know the only region with an undisputed direct Dorian heritage is Tsakonia, a place where the local language still retains may archaic Dorian elements. However, (although I am not aware of any DNA studies) their population does not appears any different to the rest of the Greeks.

kropokkuru wrote: "Though there is the fact that Scandinavia and Russia have injected small pulses of religious dissenters into Mexico in comfortably post-Columbian times, this sounds unlikely to me. Could you please share the link?"

Hammer's study of YDNA from the lower 48 US states turned up a trace of type N TAT at a very low level (0.3%) among Native Americans which was, however, much higher than the level among Europeans in the study, (0.1%). This result could be due to sampling effects or highly selective admixing, but it also is consistent with the "three major waves plus a trickle" model of migration to the Americans. (None of the Native American populations were from the Pacific Coast where the Russians had a presence.)

pconroy said...

"Fascinating, are you aware that the Irish Gaelic word for river is "Abhainn", pronounced "OW-an", and that the Welsh word for river gave us "Avon" in Stratford-on-avon... Makes is seem like the Iroquoian word is similar to the Celtic word?"

I was looking at a Proto-Celtic word list earlier this weekend, and interestingly, although the word for 'water' is akwa in Proto-Celtic, the word for 'river' is awa, which I suspect might be the archaic form.

Regarding the NE Asian admixture in Europe, it appears that there is some evidence for existence of a possible Dene-Yeniseian substrate in the Balkans. Such a substrate in Europe would have been likely if the Dene-Yeniseian link to Basque is accepted, which implies a migration from Asia through Europe, with subsequent replacement by Indo-European languges.

One common tool in the search for language substrates is the analysis of place and water names. I happened to have been looking at Thracian and Dacian when this article was posted in the blog. The words *alda (noun), *alta- (adj.) meaning 'swamp, waterlogged place' have been reconstructed from Dacian. Possible IE roots are listed as *olda, *olta ("water", "odorous"), but however these words also closely resemble 'wet' in Kott ūra and Kett ultu, both of which are Yeniseian languages. (The Basque word for 'water' is ur.) Likewise, *auras, *auro 'water, moisture, pool' with possible IE roots *uer and *au(e)r 'wet, moisten'. In Nottaway, a North American Iroquoian language, the equivalent is ya'ora.

The authors of the Wikipedia article in which these possible Dacian words are listed do note: "This is because there is no guarantee that the substratum words are, in fact, Dacian. Instead, they could derive from other, unknown or little-known tongues at some period current in Dacia or Moesia: for example, possible pre-Indo-European language(s) of the Carpathians."

'After studying the autosomal amixture data of a few Cretans and comparing them with that of Greek Cypriots, they are strikingly similar, with Cretans having only slightly higher Northern European admixture. Therefore, I would suggest that the vast majority of the Northern admixture in modern Greeks came from the Slavs and to a lesser extent from Germano-Celtic tribes.'

We don't have to take a clean pass and walk it out of the ball park. Crete as well as Cyprus are islands with pre-existing populations which have been at some point of time been colonized by Greeks and have been Hellenized afterwards. With that in mind, neither of these two island groups 'as a whole' can be considered as the epitome of Mycenean Greeks which eventually hail from mainland Greece. Crete and Cyprus were inhabited by southern peoples like Minoans, Phoenicians and pre-Greek Cypriots. If Crete is a little more northern than Cyprus as you claim, than it is likely that the original Mycenean Greeks or/and later the Dorians who invaded the islands were even more northern, given that Crete was inhabited by 'Soutern' Minoans. It would therefore make sense that mainland Greeks would be more Northern too.

High-German is Bavarian, last tribe to become germans and bavarians colonised austrian. and Austria formed in 998AD from bavarian blood.

V Robazza,

That is an oversimplification that ignores that Germanic first developed in what is now NE/NCentral Germany over 3,000 ya and thus spread from there in all directions, including S. In Roman times, an old German was already spoken along the S Rhine and between the Danube and the Alps.

There is of course some Frankish influence, in particular in what was "North Franconia" and surrounding areas. But a diaspora always retains the oldest features, while an old settled region has the most advanced changes. The only way to disentangle this seeming contradiction is that the S was mostly settled and influenced from S/ SE Central Germans, which spoke an old German for the longest time, and had developed novelties unknown in the N Central and N parts - divided by the Mittelgebirge.

"But a diaspora always retains the oldest features, while an old settled region has the most advanced changes".

That has been long accepted as the case in Polynesian languages. Hawaiian, NZ Maori and Easter Island have the oldest features. In fact the Easter Island language preserved features of Western Polynesian. Tahitian and Raratongan have changed considerably.

Eurologist, Frankish and German cannot be regarded as different branches of the Germanic tree because they are different categories. Low Frankish, colloquially referred to as Dutch, is regarded as a seperate language now, however, there are also Frankish dialects which are considered as German dialects. Some, like Ripuarian (around Cologne) are Middle German, others, like East Franconian (around northern Bavaria) are Upper German. (Middle and Upper German together are called High German, in contrast to Low German.) Actually there was no uniform Old German but different dialects, and the High German sound shift cut the Frankish dialect area into three parts. Likewise there were no Germans in antiquity and the early middle ages, but different West Germanic tribes. Some part of them eventually grew together to form the Germans, under the exclusion of formerly close relatives, like the Anglo-Saxons for instance.

As far as I know, it's not clear that the High German sound shift originated in the Bavarian dialect area. The Alemannic area may have been at the origin as well. However, while the core of the Bavarians, the Marcomanni, had been living in Bohemia for some time, the core of the Alemanni, the Semnones and Suebi, were from what is now Brandenburg and surroundings, i.e. from north of the Mittelgebirge.

Also, according to the prevalent Germanistic opinion, the High German sound shift started around 500 AD, maybe even as late as 600 AD and wasn't completed before the 9th / 10th century:

As someone who is 50% English and 50% Italian by descent, and on 23andMe is Speculatively divided between the References of "Italian", "British/Irish", "French/German", and "Iberian, I would appreciate it if someone may please better clarify what these Dodecad Results are trying to say to me. Dodecad's readings are more or less similar to those of Eurogenes for me. I'm interested in things like, as far as ancient/historical groups or populations, where I seem to pull or be more Admixed with, i.e, "Celtic", "Germanic", "Italic", etc. By the way, I am aware these are modern labels which may or may not always be too accurate, but you will know what I mean. By "Celtic", I would be meaning the "indigenous" populace of Britain, whereas "Germanic" would be the Post-Roman colonial one. By "Italic" I mean the "indigenous" populace of Italy, as opposed to outside migratory and colonial events. KIT# 213249*DODECAD V3WEST EUROPEAN-40%MEDITERRANEAN-31%WEST ASIAN-15%EAST EUROPEAN-8%SOUTHWEST ASIAN-4%#Population1 TSI-Distance: 11.82 TUSCAN-Distance: 11.863 TUSCAN-Distance: 12.034 FRENCH-Distance: 13.285 FRENCH-Distance: 13.766 N ITALIAN-Distance: 13.937 CEU-Distance: 15.358 SLOVENIAN-Distance: 15.89 N EUROPEAN-16.2410 ARGYLL-17.26Here is a different reading.*EUROGENES EUTESTATLANTIC-21%NORTH-CENTRAL EURO-21%EAST MED-15%WEST MED-15%SOUTH BALTIC-10%WEST ASIAN-8%EAST EURO-8%MIDDLE EASTERN-2%# PopulationFR-Distance: 9.07NORTH ITALIAN-Distance: 9.12AT-Distance: 10.69PT-Distance: 11SERBIAN-Distance:11.51TUSCAN-12.08WEST & CENTRAL GERMAN-Distance:12.52ES-Distance: 12.55RO-Distance: 12.74NL-Distance: 13.61Just to show a more tossed-up pic, this is my 4 Population Approximation on the EUTest1 English+GR+Scottish+Tuscan2 English+GR+IE+Tuscan3 English+GR+Orcadian+Tuscan4 DK+GR+Scottish+Tuscan5 GR+NL+Scottish+Tuscan6 DK+GR+IE+Tuscan7 GR+Orcadian+Orcadian+Tuscan8 DK+GR+Orcadian+Tuscan9 GR+IE+NL+TuscanOn nearly all the Mixed Populations, I get consistent "Tuscan", "German", "English", Cornish", "Orcadian", "French", "North Italian"-type readings in 1, 2, and 3, and it is in the 4 that "Greek" shows up. On Dodecad, "Greek" is listed in my Secondary Population Sources.

Current locations of Male Y-DNA Haplogroups can be viewed as a map of conquest. Human groups have always been in conflict, especially as weapon technology improved and animal predators were not so successful in limiting human population growth. Agriculture greatly increased human population which led to expansions - conquests.

Many of the Y-DNA Haplogroup areas are probably associated with improved weapon technologies or social organization. The first group of men who invented the sling, or spear thrower, or bow and arrow, or musket would have been able to conquer all their neighbors, replacing the males in these areas.

Among hunter gatherers their no reason to let any of the males they conquered live. After agriculture was implemented the conquering group would leave the conquered males alive to grow products to enrich the conquering group.

"Among hunter gatherers their no reason to let any of the males they conquered live".

I don't think that is altogether correct. We have many examples of regions containing a considerable variety of Y-DNA haplogroups. That situation wouldn't occur under your scenario. Although I agree that improved weapons would provide an advantage for a particular group that advantage is likely to be as much to do with hunting as destruction of other human groups. I am reasonably sure Y-DNA expansion is the result of expanding technology but that technology seems to have been shared (eventually?) with groups contacted during expansion. In fact particular technologies often expand far further than the Y-DNA that developed it.

"Among hunter gatherers their no reason to let any of the males they conquered live. After agriculture was implemented the conquering group would leave the conquered males alive to grow products to enrich the conquering group."

Let me add to terryt's response.One of the things Lewis Binford demonstrated so well was that in seasonal climates, most hunter-gatherers really don't have time or energy to run around conquering anybody. They are too busy worrying about food and shelter. When resources get scarce, there's plenty of opportunity for starving or freezing to death -- there would be plenty of chances of lineages dying out on their own -- along of course with the "Red Queen" rule about keeping ahead of pathogens -- which of course becomes more dire when the group is closely related. Lethal recessive genes abound in the human genome. And if you are going to expend energy in conflict over food, it means you are already in a tough situation.

There may have been parasitic HG "conquerers." But this kind of parasite would have been especially unsuccessful, because the energy needed would exceed the return. The stupider they were, the more likely they would resort to violence.

Just some Hard-facts: Finno-Ugric language group was born in N-Eastern Europe, until the roots of ancient IE language groups go back to Asian continent. In the Eurasiansupercontinent, there are more native speakers of IE languages in the ASIAN continent than in Europen continent. (Just remember the large IE speaking populations of India Pakistan Iran)However, the 97% of Finno-ugric speaking people live in Europe. Therefore to call finno-ugric languages as "asian languages" is laughable illogical, unscientific andmisleading.

TYPE IN ENGLISH WIKIPEDIA:"Proto Indo Europeans"http://en.wikipedia.org/wiki/Proto-Indo-Europeans

and see the maps about original ancient IE people!France Britain Italy haven't signifficant proto indoeuropean genes.The Germanic people have also very very low ratio of ancient IE haplogroup markers (R1a),only Eastern Europeans have high ratio of original proto IE haplogroups markers.Western European languages belongs to IE language group,but in very very distant way.(Have you ever heard about Language-shift? The IE linguistic effect spreaded as a lingua franca between many many populations during thousands of years)The real genetical IE people equal with the Eastern European slavic people

It looks like R1a had a lot to do with the spread of the so-called SATEM (versus CENTUM) group of IE speakers (ie., Slavic, Indo-Iranian, etc,)

If you look at the maps Dienekes posted from this article, you’ll see that two branches of R1a-M417 (R1a1a1*) -- Z282 and Z93 -- pretty nicely match the current extent of traditional SATEM IE languages.

The article also gives a R1a-M417 (R1a1a1*) coalescence at around 5000-3000 BCE. 5000 years ago matches the date for the break away of the eastern IE languages suggested by guys like Ivanov and Starostin.

The article says also that it looks like R1a (M420) started in the Middle East, around Iran. Which is on the money for Indo-Iranian, meaning that maybe where R1a speakers converted to Indo-European.

(“Based on spatial distributions and diversity patterns within the R1a-M420 clade, particularly rare basal branches detected primarily within Iran and eastern Turkey, we conclude that the initial episodes of haplogroup R1a diversification likely occurred in the vicinity of present-day Iran.”)

Actually, it seems like the parent haplogroup R had more to do with Altaic and Turkic than with IE.

Take a look at this list of I guess mainly relict steppes and Central Asian groups. Looks like most of the R and R1a are Turkic or Altaic speakers, not Indo-European.

This matches up with the observation that both R and its sister clade Q appear to have moved west together from South East Asia. And suggests that R -- just like Q -- was at the time speaking something other than Indo-European. Nobody I think connects the Q gene with IE.

“Interestingly, the monophyletic group formed by haplogroups R and Q, which make up the majority of paternal lineages in Europe, Central Asia and the Americas, represents the only subclade with K2b that is not geographically restricted to Southeast Asia and Oceania. Estimates of the interval times for the branching events between M9 and P295 point to an initial rapid diversification process of K-M526 that likely occurred in Southeast Asia, with subsequent westward expansions of the ancestors of haplogroups R and Q.”

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